CSIRO PUBLISHING www.publish.csiro.au/journals/is Invertebrate Systematic.?, 2003, 17, 15-26

Andesianidae, a new family of monotrysian moths (LepidopteraiAndesianoidea) from austral South America

Donald R. Davis ' and Patricia Gentili

^Department of Systematic Biology-Entomology, National Museum of Natural History, PO Box 37012, Smithsonian Institution, Washington, DC 20560-0127, USA. BTo whom correspondence should be addressed. Email: [email protected]

Abstract. The new family Andesianidae is proposed for the genus Andesiana, consisting of three previously described species, A. brunnea Gentili, A. lamellata Gentili and A. similis Gentili, from southern Argentina and Chile. Originally described in the ditrysian family Cossidae, Andesiana has been found to possess a monotrysian female reproductive system. The principal autapomorphies for the family are the elongate second segment of the labial palpi, presence of a male tibial pouch and associated femoral hair pencil and, within the monotrysian Heteroneura, possibly the strongly dimorphic antennae (broadly bipectinate in male, filiform in female). Other significant characters of the group include a greatly reduced haustellum, male gen i tali a with a broad vinculum, acuminate uncus, scutiform juxta and a pronounced saccular lobe of the valva that bears a subapical, setose appendage within a shallow, mesal cavity. The female genitalia possess a short cloaca and paired anal papillae and lack signa. Although known to be retained only in the female of A. lamellata, the presence of a two-branched Rs in the hindwing may indicate an origin of the Andesianidae basal to the Nepticuloidea. In contrast, the structure of the wing coupling apparatus for the family suggests a later appearance. Because no synapomorphy is known linking the Andesianidae with any other superfamily, the new superfamily, Andesianoidea, is proposed.

Introduction The monotrysian Heteroneura, a questionably mono- Field investigations conducted in the southern temperate phyletic assemblage of five distantly related superfamities, is regions of South America, largely during the decade of believed to represent the basal group of the infraorder 1979-1989, significantly increased our knowledge of the Heteroneura, as indicated by the (1) retention of a primitive more primitive, basal groups of Lepidoptera. As a direct monotrysian reproductive system (Dugdale 1974), with a result of these collective efforts, three new families of moths single reproductive opening in the female, ventral common have been documented including the Heterobathmiidae oviduct and absence of a tubular ductus seminalis; (2) (Kristensen and Nielsen 1979, 1983), Palaephatidae (Davis presence of a tergosternal connection (Kyrki 1983) 1986) and, in this report, Andesianidae. Another result of immediately caudad to the first abdominal spiracle (Fig. 23), that fieldwork was the discovery of four families previously formed by a ventrocaudal process from tergum I connecting not known to occur in either the New World (Neopseustidae, with the anterolateral extension of sternum II; and (3) often Davis 1975 and Davis and Nielsen 1980; Eriocottidae, Davis more generalised distribution of wing microtrichia. Major 1990), South America (Micropterigidae, Kristensen and synapomorphies shared with the ditrysian Heteroneura Nielsen 1982), or the southern hemisphere (Prodoxidae, include: (/) reduction of the hindwing Rs system (i.e. Nielsen and Davis 1985). heteroneurous venation, with Sc and R fused from the basal The genus Andesiana Gentili, with three included species fourth of the wing to the wing margin and Rs typically {A. brunnea Gentili, A. lamellata Gentili and A. similis unbranched); (2) frenulum-retinaculum wing coupling; (3) Gentili) from southern Argentina and Chile was proposed composite male frenulum; (4) presence of precoxal bridge originally in the ditrysian family Cossidae (Gentili 1989). (Nielsen and Kristensen 1996); and (5) loss of the first Later questioning the family placement of this genus, PG abdominal sternum (Brock 1971; Kristensen 1984). then requested DRD to examine specimens of Andesiana. Monophyly of the monotrysian Heteroneura (or Monotrysia) After dissecting a female, DRD recognised that it is inadequately supported, with two possible synapo- represented a new family of monotrysian moths. morphies proposed (Davis 1999): (7) presence of spine

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Figs 1-4. Adults of Andesiana. 1, Andesiana lamellata, male (45 mm); 2, Andesiana lamellata, male (45 mm); 3, Andesiana lamellata, female (54 mm); 4, Andesiana similis, male (35 mm). (Wingspan in parentheses). combs (pectinifers) on the male valva (absent in whose larvae are known to be stem borers. Consequently, Tischerioidea, Palaephatoidea and Andesiana); and (2) adult Andesiana may be easily mistaken for a member of the complete or partial subdivision of the second abdominal ditrysian family Cossidae, unless the female internal sternum into a smaller anterior sclerite (S2a) and a usually reproductive morphology is examined. Although their much larger caudal sclerite. The latter condition is often biology is unknown, the greasy condition, which develops in associated with the development of a pair of oval areas many museum specimens, as well as their relative body size, (windows) that largely separate the more sclerotised strongly suggests that larval Andesiana are also stem borers. anterior/posterior regions of sternum II. Lightly sclerotised Adult Andesianidae are phototrophic and are readily windows also occur on S2 in some ditrysian families (e.g. attracted to ultraviolet light. Acrolophidae). Partial to complete subdivision of sternum II, Andesianidae is characterised by at least two often associated with paired sternal windows, is known to autapomorphies: (7) elongate second segment of the labial exist in other primitive glossatan families (e.g. Eriocraniidae palpi (Fig. 5) and (2) presence of male tibial pouch and and Acanthopteroctetidae, Davis 1978; Lophocoronidae, associated femoral hair pencil (Fig. la, b). The second labial Nielsen and Kristensen 1996) and may be part of the palpal segment is nearly 4 % the length of the apical segment. Lepidoptera Glossata groundplan. The latter terminates in an elongate sensory pit (organ of Adult Andesiana are moderately large, heavy bodied, vom Rath; Figs 5, 14), which often is found everted in dried cossoid-appearing moths quite unlike any previously specimens (Fig. 12). Specialised organs for pheromone recognised member of the Monotrysia Heteroneura. Their dispersion are sparingly developed among the monotrysian wing span ranges from 27 to 61 mm (1.19-2.38 inches), with Heteroneura, with the notable exception of Palaephatidae a maximum size nearly twice that of the largest previously (Davis 1986) and one unusual occurrence within the known monotrysian species (e.g. 32 mm in female Incurvarioidea (Wojtusiak 1999). The males of Andesiana Palaephatus dimorphus Davis). Moreover, the bodies and possess a rather elaborate androconial organ on the hind leg wings of dried specimens often become greasy, a condition consisting of an elongate hair pencil composed of numerous typically occurring in other families, particularly Cossidae, filamentous setae that arise subapically from the femur and The new monotrysian moth family Andesianidae Invertebrate Systematics 17

hf \ S= R1 Rs1

Figs 5-7. Adult morphology of Andesiana. Andesiana lamellatu 5-7a: 5, frontal view of head (1.0 mm); 6, female wing venation; 7, male legs (5.0 mm); 7a, detail of male femoral hair pencil and tibial pouch (0.5 mm). 7b, Andesiana similis: male femoral hair pencil and inflated tibial pouch (1.0 mm). (HP = hair pencil; H - haustellum; IS = intercalary sclerite; OR = organ of vom Rath; TP = tibial pouch; scale lengths in parentheses). extend approximately 1/2-2/3 the length of the tibia. The external wall of the pouch is densely covered with minute, distal fourth to one third of the hair pencil normally rests non-deciduous, piliform scales. Hydrostatic pressure within inside an inverted pocket within a large, membranous, the pouch everts the pocket, thereby expelling the hair pencil. inflatable pouch at the base of the hind tibia (Fig. la). The Once expelled, it is unlikely that the hair pencil can ever be 18 Invertebrate Systematics D. R. Davis and P. Gentili

Figs 8-13. Andesiana lamellata, head morphology. 8, Female antenna, near basal third of flagellum, top of photo = dorsal. 9, Detail of sensillum coeloconicum (centre) from Fig. 8. 10, Male antenna! pectinations, dorsal view. 11, Venter of Fig. 10. 12, Labial palpal segment III with subapical organ of vom Rath exerted. 13, Detail of apical sensillac in Fig. 12.

reinserted into the tibial sac. The length of the fully inflated dense scattering of long sensillae chaetica (Fig. 11), which pouch ranges from approximately half the length of the tibia are more dispersed over the female antenna (Fig. 8). in A. lamellata to nearly its entire length in A. similis (Fig. Sensillae coeloconica are scattered along the lower surfaces lb). No males of A. brunnea were available for study. of the male rami and are denser ventrally along the female Strongly dimorphic antennae (broadly bipectinate in flagellomeres (Figs 8, 9). A broadly triangular intercalary male, filiform in female) (Figs 8, 10, 11) may constitute sclerite is present between the posterioventral margins of the another autapomorphy for Andesiana. Within the scape and pedicel. monotrysian Heteroneura, development of bipectinate The forewings of Andesiana are moderately broad antennae was known to occur only in the males of some with a L/W ratio of c. 2.5 and a hindwing ratio of species of Incurvariidae (e.g. Incurvaria masculella (D. & nearly 2.0. Microtrichia are primarily restricted to the S.)). The pectinations are more developed in the males of forewing anal region. The male retinaculum consists of a Andesiana than those of Incurvaria, with the longest rami broadly triangular, subcostal flap with a densely scaled, approximately 4.5-5.0 x the length of the flagellomere. The revolute apex into which the single, large, compound dorsum of the antenna in both sexes is somewhat thinly frenulum inserts. The female frenulum is composed of covered from the scape nearly to the apex with slender, 5-6 smaller bristles arising in a separated series from bi-tridentate scales arranged in indefinite, scattered rows the base of the hindwing costal margin, thus resembling (Figs 8, 10). The male rami are naked ventrally except for a that found in certain other monotrysian genera such as The new monotrysian moth family Andesianidac Invertebrate Systematics 19

Palaephatus. Concentration of the female frenular anus and oviporus well separated and exposed. Attached to bristles near the base of the hindwing may indicate an the anteriolateral walls of the cloaca of Andesiana near the appearance of the Andesianidac later than the junction of the rectum and vagina are a pair of tendon-like Nepticuloidea, although the enlarged series present in apodemes (CIA; Fig. 41). These may be homologous to the female Nepticuloidea may also represent a derived state cloacal apodemes present in Adelidae (Davis 1998, (Davis 1986). fig. 6.8G). The hindwing radial system of female Andesiana Because no synapomorphy was found linking the lamellata is unusual among the Heteroneura in possessing Andesianidac with any other superfamily, a new superfamily, two separate branches of the radial sector vein (Rsl, 2) Andesianoidea, is proposed, pending a molecular analysis of arising near the apex of the discal cell (Fig.6). No females of the monotrysian families. A. similis or A. brunnea were available for this study. The normal condition within the Lepidoptera Heteroneura is for the radial sector to be unbranched (Scoble 1992). The extra Andesianidae, new family Rs vein reported in a few more advanced heteroneuran Type genus: Andesiana Gentili, 1989. genera (Busck 1914; Braun 1933) probably represents secondary reversals without phylogenetic significance Description (Nielsen and Kristensen 1996). Although Rs2 has been Head (Fig. 5). Vestiture of frons and vertex rough, observed only in the females of one member of the family, consisting of dense, long, piliform scales forming a the presence of Rs2 in the hindwing of the monotrysian longitudinal crest over vertex. Chaetosemata absent. Andesiana may indicate this group's origin basal to the Cranium with a pair of broad, slightly raised swellings at Nepticuloidea. This contradicts the somewhat weaker vertex. Compound eyes naked, large, with interocular index evidence based on a comparison of female frenular structure. of ~1.25 and eye index of ~0.93; all species nocturnal and Sternite IX (vinculum) of male Andesiana (Figs 24, 29, attracted to light. Lateral ocelli absent. Antenna long, -0.55 34) resembles that of several monotrysian genera of length of forewing in both sexes, with 66-93 segments in Incurvarioidea and Nepticulidae in being typically long and males, 64-88 in females; scape cupuliform, covered by white broad. The valvae are unique in being almost completely scales, without pecten; intercalary sclerite broadly divided, with a prominent, digitate, saccular lobe bearing a triangular; pedicel a slightly globose cylinder; flagellomeres small, elliptical, strongly setose appendage (ASL; Fig. 33) cylindrical, with two long pectinations (rami) in male; rami within a subapical cavity, which opens mesally (Fig. 33). The arising ventrally from flagellomere, near proximal end; a juxta in Andesianidac consists of a sclerotised, well slightly swollen area bearing a concentration of sensillae developed, roughly quadrate to medially constricted plate chaetica located distal to origin of rami. Tentorium with well more similar to that present in some Incurvarioidea (Davis developed anterior arms, decreasing in diameter posteriorly; 1999). A juxta is poorly developed to absent in the dorsal arms absent. Labrum reduced to a shortened, broadly Nepticuloidea, Tischerioidea and Palaephatoidea. triangular sclerite; pilifers indistinct. Mandibles vestigial, The female reproductive system of Andesiana does not mostly membranous, with only a narrow, sclerotised lateral agree closely with any known monotrysian family. The rim preserved. Haustellum reduced, approximately equal in Incurvarioidea are typically distinguished by their greatly length to first segment of labial palpus; food channel extensible, piercing ovipositor, elongate posterior shallow, with a row of minute, slender legulae bordering apophyses, extended cloaca and enclosure of the eighth channel. Maxillary palpus well developed, 5-segmented; segment normally within the greatly enlarged seventh. length ratio of segments from base: -1:1:1.6:2.5:0.5. Although differing less in general morphology from the Labial palpus 3-segmented, with extremely elongate second Palaephatidae, female Andesiana possess paired anal segment; length ratio from base for males is 1 :3.5 : 1.08; for papillae (not developed in Palaephatidae) and lack a females, 1:6.3:1.2; apical segment with elongate, eversible terminal, median sensory ridge (a possible synapomorphy sensory pit (organ of vom Rath), -0.33-0.50 length of apical for Palaephatidae and Tischeriidae). Tergum IX + X of segment. Opostegidae varies in either lacking anal papillae, or Thorax. Wings broad; forewing almost rectangular. possessing one- or, usually, two-lobed papillae (Davis 1989). Wingspan: 27.3^15.0 mm in males, 47.4-61 mm in females. Paired anal papillae also occur in Tischeriidae, in close Radial system of forewing five-branched, with R arising association with a median sensory ridge (Davis 1986). The from basal third of discal cell, and branches of radial sector caudal terminations of the alimentary tract (anus) and from the areola (accessory cell). Dorsal vestiture of forewing reproductive tract (oviporus) in Andesianidac merge within smooth except for a patch of more erect scales at end of a broad, poorly defined cloaca consisting largely of a discal cell, which are also present but less dense on surrounding membranous fringe which opens terminally on hindwing. Microtrichiae generally absent over both wings segment X. A cloaca is absent in Palaephatidae, with the except for anal patch near the base of forewing. Retinaculum 20 Invertebrate Systematics D. R. Davis and P. Gentili

Figs 14-19. Andesiana lamellate, adult morphology. 14, Labial palpal segment III showing aperture of organ of vom Rath. 15, Apical sensillae of maxillary palpal segment V. 16, Lateral view of prothoracic pretarsus. / 7, Dorsal view of Fig. 16 showing pseudempodial seta (centre). 18, Ventral view of Fig. 16. 19, Detail of unguitractor plate from Fig. 18. a broad ventral lobe from costal margin in male, with curved, ventral surface; structure of pretarsus typical for most densely scaled apex; retinaculum in female consisting of stiff Lepidoptera families; ungues long, strongly curved; arolium, scales along costal margin. Hindwing with veins Sc and R pseudempodial seta, and median unguitractor plate well fused from basal fourth to apex; Rs unbranched except with developed (Figs 16-19); pulvillae triangular, densely setose, Rsl and Rs2 preserved in female Andesiana lamellata somewhat reduced, less than length of ungues. Prothorax (Fig. 6). Base of medial vein present and forked within discal with lateral cervical sclerite evenly curved as in cell. Medial veins always arising separately from cell. Crinopterigidae (Davis 1999); lateral angle produced as thin 1A+2A forked over basal third. Male frenulum a single, plate with thickened ventral margin (Fig. 20). composite spine; female frenulum consisting of 4-6 smaller, Metafurcasternum with furcal arms broad, tapering, and free; well spaced spines along costal margin. Legs elongate anteromedial process relatively short; apex of mesal lamella (Fig. 7); tibial spur formula 0-2^1; spurs asymmetrical, broadly rounded (Fig. 21,21a). densely covered by short hairs, cylindrical, with acute apex. Abdomen. Sternum 1 absent; sternum 2 subdivided, Epiphysis flattened, elongated, 0.56-0.60 length of fore tibia with a pair of oval, lightly sclerotised windows between S2a in male, 0.48-0.52 in female. Hind tibia of male with a basal, and S2; S2a broadly U-shaped, less than half width of S2 eversible pouch, which at rest holds tip of elongate hair (Fig. 22). Tergosternal connection present caudad to first pencil arising from inner subapical surface of hind femur abdominal spiracle. Coremata absent in male; corythrogyne (Fig. la, b); hair pencil and pouch absent in female. Tarsus absent in female. Spiracle 8 well developed and functional in and apical half of tibia with numerous scattered spines along female, vestigial in male. Eighth sternum of male widely The new monotrysian moth family Andesianidae Invertebrate Systematics 21

SAI

21a

Figs 20-23. Andesiana lamellata, thoracic morphology. 20, Anterior view of prothoracic lateral cervical sclerites (0.5 mm). 21, Posterior view of metathorax (1.0 mm); (a), lateral view of metafurcasternum (1.0 mm). 22, Ventral view of second abdominal sternum (2.0 mm). 23, Lateral view of abdominal base (2.0 mm). (APM = anteriomedial process of metafurcasternum; FA = furcal apophysis; IL = intercoxal lamella of basisternum; LP = laterophragma; ML = mesal lamella of metafurcasternum; SAF = secondary arms of metafurcasternum; S2a = anterior sclerite of second sternum; Scm = scutellum; 1 SP = first abdominal spiracle; T2 = second tergum; TP = tubcrculate plate; TsC = tergosternal connection; WP = wart-like protuberence; scale lengths in parentheses). excavated along caudal margin to accommodate broad triangular, fused to tegumen but with suture moderately vinculum; sternum VIII of female densely setose, caudal distinct; apex of uncus acuminate, slightly curved ventrad, margin either entire and narrowly rounded or with median with a double row of short, stout spines. Gnathos absent. cleft (in A. similis). Socii reduced, present as small, membranous, setose pads Male genitalia (Figs 24-38). Ninth segment a ventrolateral to tegumen. Valva strongly fused basally by a continuous ring consisting of an elongate, broad, roughly broad, strongly developed transtilla; valva divided for nearly quadrate sternum (vinculum) and much more narrow tergum its entire length into a broad cucullus and stout, digitiform, (tegumen); anterior half of vinculum slightly flared, with a sacculus (Figs 28, 33); mesal subapex of saccular lobe sinuate to concave anterior margin (Figs 24, 29, 34); lateral shallowly excavate, containing an elliptical, densely setose angles of tegumen with variously developed, paired appendage (ASL; Fig. 33); pedicel of sacculus either with a processes (Figs 25, 30, 35). Tenth tergum (uncus) variably membranous, mesal groove extending from base to subapical 22 Invertebrate Systematics D. R. Davis and P. Gentili

Figs 24-38. Male genitalia and female abdominal segments 8-10 of Andesiana. 24-28, Andesiana lamellata: 24, ventral view, male genitalia (1.0 mm); 25, dorsal view of tegumen and uncus; 26, aedeagus; 27, juxta; 28, mesal view of valva (1.0 mm). 29-33, Andesiana similis: 29, ventral view (1.0 mm); 30, dorsal view of tegumen and uncus; 31, aedeagus; 32, juxta; 33, mesal view of valva (1.0 mm). 34-36, Andesiana brunnea: 34, ventral view (1.0 mm); 35, dorsal view of tegumen and uncus; 36, aedeagus. 37-38, Andesiana similis: abdominal segments 8-10 of female: 37, lateral view (0.5 mm); 38, ventral view. (ASL = appendage of saccular lobe; PAn = papilla analis; S8 = eighth sternum; So = socii; T8 = eighth tergum: Tn = tegumen; Tr = transtilla; U = uncus; scale lengths in parentheses).

cavity (in A. lamellata, A. similis) or completely sclerotised enlargement evident in one species {A. brunnea); cornuti of and without groove (in A. brunnea). Juxta a short, vesica either scattered or arranged in two longitudinal rows. sclerotised, subrectangular plate, moderately to strongly Female genitalia (Figs 37^41). Ovipositor non- constricted at middle (Figs 27, 32); distal end ventrally piercing, short, largely enclosed within eighth abdominal attached to aedeagus through anellar membrane. Anellus segment. Segments IX + X fused, represented primarily by a with a pair of elongate, laterodorsal concentrations of small pair of small, setose anal papillae immediately dorsal to spines. Aedeagus moderately stout and short, -equal to valva terminal cloacal opening. Posterior apophyses slender, in length; coecum poorly developed, only a slight basal approximately equal to eighth segment in length. Anterior The new monotrysian moth family Andesianidae Invertebrate Systematics 21

Figs 39—41. Female gcnitalia of Andesiana lamellata. 39, Parasagittal view of abdominal segments 7—10 (1.0 mm). 40, Ventral view of abdominal segments 8-10 (1.0 mm). 41, Dorsal view of Fig. 40, with tergum removed (1.0 mm). (AA = anterior apophysis; BC = bursae copulatrix; CG = duct of colleterial gland; CIA = cloacal apodeme; CO = common oviduct; Cl = cloaca; DB = ductus bursae; DS = ductus seminalis; PA = posterior apophysis; PAn = Papillae analis; R = rectum; RR = rectal reservoir; SP = spermathecal papilla; Sp8 = eighth abdominal spiracle; T7 = seventh tergum; T8 = eighth tergum; Va = vagina; scale lengths in parentheses).

apophyses arising from eighth sternum, -1.6 % length of oviduct, membranous, without signa. Spermatheca with posterior apophyses. Cloaca broad, extremely short, formed papilla greatly enlarged, joining vagina anterior to junction primarily by membranous fringe surrounding cloacal of common oviduct. Vagina broad at anterior junction with opening. A pair of cloacal apodemes (CIA; Fig. 41) arising spermathecal papilla, then narrowing beyond junction with from anteriolateral walls of cloaca. Common oviduct oviduct. Common duct of colleterial gland joining vagina entering vagina ventrally. Bursa copulatrix located dorsal to dorsal ly, caudad to spermathecal papillae. 24 Invertebrate Systematics D. R. Davis and P. Gentili

Andesiana Gentili, 1989 Type species: Andesiana lamellata Gentili, 1989: 21, original designation.

Description Adult As described for the family.

Species synopses

Andesiana lamellata Gentili, 1989

Description Q

Adult (Figs 1-3) Length of forewing: S 15.7-26.1 mm, ? 21.5-26.1 mm. Largest of the three species, with the most evident sexual dimorphism: females often 1/3 larger than males. Vestiture of male forewing grey with a metallic sheen, covered by a reticulate pattern of reddish-brown; markings more dense and forming three irregular patches (sub-basal, median and lamellata sub-terminal) along costa, and another patch touching hind similis margin in cubito-anal area. A well defined bar of darker ^fcbrunnea brown composed of erect, dense scales at end of discal cell. Hindwings grey with a diffuse reticulated brownish pattern, lighter towards wing base; a darker bar along end of discal J&_^t( cell as in the forewing, but not as prominent. Females with a less pronounced wing pattern. Vestiture of the head, thorax and abdomen a combination of piliform scales and broader Fig. 42. Distribution of Andesiana lamellata, A. similis and A. scales forming crests and tufts on head, collar and thorax; brunnea in southern Chile and Argentina. abdomen grey. Male genitalia (Figs 24-28). Laterodorsal processes of tegumen triangular, short, acute. Transtilla a broad plate tapering slightly anteriorly, with more narrow, free anterior Rio Negro) and Chile (Malleco and Valdivia) between end bilobed. Juxta strongly constricted at middle; anterior 38°40' and 42°S. Adults fly during the spring into early margin smoothly rounded. Valva with apex of cucullus more summer (late September to mid January, with isolated extended than in other species; digitiform saccular lobe of records in February and April) in Nothofagus pumilio and N. valva relatively elongate, subapical cavity elliptical. antarctica forests. They occur over a broad altitudinal range, Aedeagus with cornuti relatively reduced in size, numerous between 100 and 1750 m. and scattered; anellar spines in two, relatively broad, lateral patches. Female genitalia (Figs 39—41). Anal papillae lightly Remarks sclerotised, posterior apophyses reaching the anterior margin Of the other two species in the genus, only A. similis is likely of eighth tergum. Posterior margin of sternum VIII arcuate, to be confused with A. lamellata. The male genitalia of A. sclerotised. similis are characterised by long, curved, digitate, laterodorsal processes of the tegumen (short in A. lamellata); dorsal plate of transtilla sub-trapezoidal (rectangular in A. Distribution (Fig. 42) lamellata); juxta basally expanded, with expansions Andesiana lamellata is the most common species of the terminating in acute apices (not expanded, more simple in^4. group, with a geographical range in Argentina (Neuquen and lamellata). The new monotrysian moth family Andesianidae Invertebrate Systematics 25

Andesiana similis Gentili, 1989 defined, dark brown bar composed of erect dense scales at end of discal cell. Hindwings are almost denuded of scales, Description with scattered brown scales along margins; a darker bar Adult (Fig. 4) along end of discal cell as in forewing. Vestiture of head, Length of forewing: S 17.4-18.2 mm, 2 21.15 mm (only thorax and abdomen with same combination of crests and one female known). Vestiture of male forewing steel-grey tufts, but more uniform in colour and dark reddish brown as with a metallic sheen; pattern as in A. lamellata but not as in wings. prominent and dominated by a white subapical band oblique Male genitalia (Figs 34-36). Laterodorsal processes of across wing; also a series of dark spots at terminus of veins. tegumen reduced to a pair of short denticles. Transtilla (not Hindwings whitish-grey with copper iridescence on veins. illustrated in Fig. 34) broad, tapering to more narrow anterior Female with less pronounced wing pattern. Vestiture of head, margin. Juxta constricted at middle, anterior margin thorax and abdomen as in A. lamellata. rounded. Saccular lobe without longitudinal groove. Male genitalia (Figs 29-33). Laterodorsal processes of Aedeagus short, vesica with a conspicuous subapical tegumen long and curved towards uncus. Transtilla broad, sclerotised plate and two small patches of cornuti; anellar strongly tapered anteriorly; anterior margin superficially spine patches similar to A. lamellata. sinuate. Lateral margins of juxta almost parallel, proximally Female genitalia. Unknown. expanded; anterior margin of juxta irregular. Valva with cucullar margin more truncate than in A. lamellata; Distribution (Fig. 42) digitiform saccular lobe of valva relatively shorter and more Andesiana brunnea is known from a single specimen from clavate than in A. lamellata; apical cavity more circular. Argentina (Isla Victoria, Lago Nahuel Huapi, Neuquen) Aedeagus with cornuti more elongate and arranged in two collected in early October. subapical rows; anellar spines arranged in more slender lateral patches than in other species. Remarks Female genitalia (Figs 37-38). Anal papillae The small size, dark brown colouration, and more slender sclerotised. Posterior apophyses slightly surpassing anterior wings of this species easily distinguish A. brunnea from the margin of eighth tergum. Eighth sternum sclerotised over foregoing species. The male genitalia possess short, paired, distal half; distal margin protruding, with a strong median laterodorsal processes on the tegumen more similar to those indentation. of A. lamellata and distinct from the elongate processes of A. similis. Distribution (Fig. 42) Andesiana similis commonly occurs in Nothofagus dombeyi Acknowledgments forests with an understory of Chusquea culeou in Argentina We dedicate this paper to the memory of Ebbe S. Nielsen, (Neuquen) around 40°S. Adults fly during the spring (late whose interest in the lower Lepidoptera was both inspiring October to mid December). Their altitudinal range is more and broad in scope. It is interesting to note that one of the last restricted, between 640 and 950 m. discussions DRD had with Ebbe centred on the relationships Remarks of Andesiana, wherein he queried 'was Andesiana actually a monotrysian?' We hope this report answers his question. This species can only be confused with A. lamellata, from From the Department of Systematic Biology, Smithsonian which it differs by its smaller size and lighter colouration, Institution, we thank Vichai Malikul and Young Sohn for the particularly by the sub-apical white band across the line illustrations, and George Venable for assistance with forewing. Andesiana lamellata also lacks the brown spots at graphics. Susann Braden of the Smithsonian Scanning the end of the veins. Differences in genitalic morphology are Electron Microscope Laboratory and John Steiner of the summarised above under A. lamellata. Diagnostic Smithsonian Photographic Services assisted with the comparison with A. brunnea is provided below. photographs.

References Andesiana brunnea Gentili, 1989 Braun, A. F. (1933). Pupal tracheation and imaginal venation in Microlepidoptera. Transactions of the American Entomological Description .Sbciefy 59, 229-268. Adult Brock, J. P. (1971). A contribution towards an understanding of the morphology and phytogeny of the ditrysian Lepidoptera. Journal of Length of forewing: 6, 12.4 mm, only one specimen Natural History 5, 29-102. known. The smallest of the three species, with generally dark Busck, A. (1914). On the classification of the Microlepidoptera. reddish-brown vestiture. Forewing pattern reduced to a well Proceedings of the Entomological Society of Washington 16,46—54. 26 Invertebrate Systematics D. R. Davis and P. Gentili

Davis, D. R. (1975). Systematics and zoogeography of the family Kristensen, N. P. (1984). Studies on the morphology and systematics of Neopseustidae with a proposal of a new superfamily (Lepidoptcra: primitive Lepidoptera (Insecta). Steenstrupia 10, 141-191. Neopseustoidea). Smithsonian Contributions to Zoology 210, 1—44. Kristensen, N. P., and Nielsen, E. S. (1979). A new subfamily of Davis, D. R. (1978). A revision of the North American moths of the micropterigid moths from South America. A contribution to the superfamily Eriocranioidea with the proposal of a new family, morphology and phylogeny of the Micropterigidae, with a generic Acanthopteroctetidae (Lepidoptera). Smithsonian Contributions to catalogue of the family (Lepidoptera: Zeugloptera). Steenstrupia 5, Zoology 251, 1-131. 69-147. Davis, D. R. (1986). A new family of monotrysian moths from austral Kristensen, N. P., and Nielsen, E. S. (1982). South American South America (Lepidoptcra: Palaephatidae), with a phylogenetic micropterigid moths: two new genera of the Sabatinca-group review of the Monotrysia. Smithsonian Contributions to Zoology (Lepidoptera: Micropterigidae). Entomologica scandinavica 13, 434, 1-202. 513-529. Davis, 0. R. (1989). Generic revision of the Opostegidae, with a Kristensen, N. P., and Nielsen, E. S. (1983). The Heterohathmia life synoptic catalogue of the World's species (Lepidoptera: history elucidated: immature stages contradict assignment to Nepticuloidea). Smithsonian Contributions to Zoology 478, 1-97. suborder Zeugloptera (Insecta, Lepidoptera). Zeitschrift fiir Davis, D. R. (1990). Neotropical Microlepidoptera XXIII. First report Zoologische Systematik und Evolutionsforschung 21, 101-124. of the family Eriocottidae from the New World, with descriptions of Kyrki, J. (1983). Adult abdominal sternum II in ditrysian tineoid new taxa. Proceedings of the Entomological Society of Washington superfamities - morphology and phylogenetic significance 92, 1-35. (Lepidoptera). Annales Entomologici Fennici 49, 89-94. Davis, 0. R. (1998). The Monotrysian Heteroneura. In 'Handbook of Nielsen, E. S., and Davis, 0. R. (1985). The first southern hemisphere Zoology, Vol. IV, Part 35. Lepidoptera, Moths and Butterflies. prodoxid and the phylogeny of the Incurvarioidea (Lepidoptera). Volume 1: Evolution, Systematics, and Biogeography'. (Ed. N. P. Systematic Entomology 10, 307-322. Kristensen.) pp. 65-90. (Walter de Gruyter: Berlin, Germany.) Nielsen, E. S., and Kristensen, N. P. (1996). The Australian moth family Davis, D. R., and Nielsen, E. S. (1980). Description of a new genus and Lophocoronidae and the basal phylogeny of the Lepidoptera- two new species of Neopseustidae from South America, with Glossata. Invertebrate Taxonomy 10, 1199-1302. discussion of phytogeny and biological observations. Steenstrupia Scoble, M. J. (1992). 'The Lepidoptera, Form, Function, and Diversity.' 6, 253-289. (Oxford University Press: New York, USA.) Dugdale, J. S. (1974). Female genital configuration in the classification Wojtusiak, J. (1999). A new type of scent organ in Lepidoptera: a giant of Lepidoptera. New Zealand.Journal of Zoology 1, 127-146. thoracic pheromone-disseminating structure in a male incurvarioid Gentili, P. (1989). Revision sistematica de los Cossidae (Lep.) de la moth (Lepidoptera, Adelidae). Journal of Natural History 33, Patagonia Andina. Revista de la Sociedad Entomologica Argentina 819-824. 45, 3-76.

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